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As Gilbert White, Darwin, and others observed long ago, all species appear
to have the innate capacity to increase their numbers from generation to
generation. The task for ecologists is to untangle the environmental and
biological factors that hold this intrinsic capacity for population growth in
check over the long run. The great variety of dynamic behaviors exhibited by
different population makes this task more difficult: some populations remain
roughly constant from year to year; others exhibit regular cycles of abundance
and scarcity; still others vary wildly, with outbreaks and crashes that are in
some cases plainly correlated with the weather, and in other cases not.
To impose some order on this kaleidoscope of patterns, one school of
thought proposes dividing populations into two groups. These ecologists posit
that the relatively steady populations have “density-dependent” growth
parameters; that is, rates of birth, death, and migration which depend strongly
on population density. The highly varying populations have “density-independent”
growth parameters, with vital rates buffeted by environmental events; these
rates fluctuate in a way that is wholly independent of population density.
This dichotomy has its uses, but it can cause problems if taken too
literally. For one thing, no population can be driven entirely by
density-independent factors all the time. No matter how severely or
unpredictably birth, death, and migration rates may be fluctuating around their
long-term averages, if there were no density-dependent effects, the population
would, in the long run, either increase or decrease without bound (barring a
miracle by which gains and losses canceled exactly). Put another way, it may be
that on average 99 percent of all deaths in a population arise from
density-independent causes, and only one percent from factors varying with
density. The factors making up the one percent may seem unimportant, and their
cause may be correspondingly hard to determine. Yet, whether recognized or not,
they will usually determine the long-term average population density.
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